A new combination in (Compositae: ), and a synopsis of the

JOSE M. BONIFACINO1, GISELA SANCHO2, AND EDUARDO H. MARCHESI3

'Laboratorio de Botinica, Facultad de Agronomia, Casilla de Correos 1238, Montevideo, ; e-mail: bonifacinoj @fagro.edu.uy 2 Divisi6n Plantas Vasculares, Museo de La Plata, Paseo del Bosque s. n., La Plata, 1900, Buenos Aires, ; e-mail: [email protected] 3 Laboratorio de Botinica, Facultad de Agronomia, Casilla de Correos 1238, Montevideo, Uruguay; e-mail: [email protected]

Abstract. A new combination, Asteropsis megapotamica, is made after studying specimens at Paris (P). A taxonomic revision of Asteropsis, with detailed descriptions, drawings, and images of A. megapotamica is presented. A lectotype for Neja macrocephala is designated. A discussion about generic relationships of Asteropsis and closely related genera is provided.

Key Words: Asteropsis megapotamica, new combination, , Podocominae, Uruguay.

The rare and geographically restricted toire Naturelle (P) described by Sprengel monotypic genus Asteropsis Less. (Composi- (1826), it has become evident to us that tae: Astereae) is endemic to southern Brazil Aster megapotamica Spreng. refers to the and northern Uruguay, and was originally same taxonomic entity described by Lessing described, along with its sole species Aster- under Asteropsis. Therefore, both names opsis macrocephala Less., by Lessing (1832). should be considered heterotypic synonyms, Independently, and based on different with Sprengel's name having priority over material from , Hooker Lessing's. and Amrnott (1836) published Podopappus Here, we present a new combination for tomentosus Hook. & Amrn. to refer to the Asteropsis and provide its taxonomic revision same taxon coined by Lessing as Asteropsis with detailed descriptions, drawings, and macrocephala. Later on, Bentham (1873) images of its only species, A. megapotamica. placed Asteropsis in the synonymy of Podo- coma Cass.; however, he did not make the Material and methods combination. It was up to Herter (1930) to publish the combination macro- Data is derived from the study of herbar- cephala (Less.) Herter, as suggested by ium specimens from BM, K, LP, MVFA, P, Bentham. SI, and US, and from field observations. For Nevertheless, in his treatment of Brazil- light microscopic examination, floral and ian Astereae, Baker (1882) recognized vegetative parts were re-hydrated and stained Asteropsis as a distinct genus, just as more in 2% safranin. Involucre shape and measure- recent authors do (Nesom, 1994a; Sancho et ments were made on live specimens. Draw- al., 2006; Nesom & Robinson, 2007; Sancho ings were made by the authors using a Wild & Karaman-Castro, in press; Brouillet et al., M3Z stereo microscope and a Leitz SM Lux in press). After having had access to type microscope with camera lucida. Terminology material of South American Astereae from follows Harris and Harris (1994), Ramayya the Herbarium of Mus6um National d'His- (1962), and Steam (1995).

Brittonia, 61(1), 2009, pp. 1-7 ISSUED: 2 March 2009 A 2008, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

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Generic relationships generic relationships of Asteropsis sug- gested by Nesom and Robinson (2007). In After its original description, Asteropsis was recent studies, Asteropsis forms an assem- sunken into synonymy with Podocoma by blage with Inulopsis, Microgyne, and Som- Bentham (1873). This was followed by merfeltia, and among those genera, the several authors including Zhang and Bremer closest relationship of Asteropsis is with (1993) who incorporated Podocoma into their Inulopsis (Sancho & Karaman-Castro, in Hinterhubera group (i.e., Blakiella Cuatrec., press; Brouillet et al., in press). Close Flosmutisia Cuatrec., Hinterhubera Sch. Bip., phylogenetic relationships between Podo- Microgyne Less., and Westoniella Cuatrec.). coma and Asteropsis were not demonstrated These authors based their inclusion of in these phylogenetic studies. Morphologi- Podocoma on the presence of beaked cypse- cally, Asteropsis differs from Podocoma by lae, a character that, of the five genera in the its densely leafy stems, solitary, strongly group, was only shared with Blakiella. radiate capitula, and cypselae that are atten- Bremer (1994) followed Zhang and Bremer uate towards the apex (vs. sparsely leafy (1993) in regards to the relationships of stems, corymbiform capitulescences of few Podocoma. However, recent morphological heads, sub-radiate capitula, and truly rostrate and molecular evidence (Karaman, 2006; cypselae in Podocoma). Nesom & Robinson, 2007; Sancho & Among the South American genera of Karaman-Castro, in press) have shown that Podocominae, Asteropsis, Inulopsis, Micro- the beaked cypselae of Blakiella and gyne, and Sommerfeltia share radiate capitula, Podocoma originated independently. usually sterile, non-rostrate, disk cypselae, Nesom (1994a) regarded Asteropsis as a and a pappus of (1)2 or 3 series of bristles. distinct genus. He also provided the first Morphologically, Inulopsis shares with Aster- detailed synonymy for this genus, accounting opsis glandular cypselae with an attenuate for most of the names ascribed to A. macro- apex (abruptly attenuate in Inulopsis and cephala, and commented on its putative gradually attenuate in Asteropsis). Asteropsis phylogenetic relationships, naming Podo- and two species of Inulopsis (I. scaposa coma, Microgyne, Sommerfeltia Less., Inu- (DC.) O. Hoffm., I. stenophylla Dus~n) also lopsis (DC.) O. Hoffim., Rhabdanthus G. L. share a pappus with an outermost series of Nesom, Laennecia Cass., and Blakiella as its very short bristles. Inulopsis differs from closest relatives. When the subtribe Podoco- Asteropsis by its single-seriate ray florets minae was described by Nesom (1994b), he with relatively wide corolla limbs (vs. 3- or included Asteropsis, other South American 4-seriate ray florets with narrow corolla limbs genera such as Blakiella, Inulopsis, Laennecia, in Asteropsis). Podocoma, and Sommerfeltia, and a group of We follow Sancho and Karaman-Castro (in Australasian genera. The genus Blakiella was press) and Brouillet et al. (in press) in later transferred to the subtribe Hinterhuberinae considering three genera of the current Podo- (Nesom & Robinson, 2007). cominae (Inulopsis, Microgyne, and Sommer- Morphological and molecular phyloge- feltia) as most closely related to Asteropsis, netic studies partially agree with the and present a key to differentiate these genera.

Key to Asteropsis and related genera

1. unbranched or only branched at the base; leaves linear, narrowly elliptic or obovate to widely obovate with margins mostly entire (lowermost leaves with lobed margins in Asteropsis); cypselae attenuate at apex. 2. Ray florets in 1 (2) series, ray corollas with relatively wide limbs, (1-)1.5-2.5 mm wide; cypselae abruptly attenuate at the apex ...... Inulopsis 2. Ray florets in 3 or 4 series, ray corollas with relatively narrow limbs, up to 1 mm wide; cypselae gradually attenuated towards the apex Asteropsis 1. Plants shrubby, fully branched; leaves elliptic (in outline) with pinnatifid margins; cypsela apex truncate or with confluent margins. 3. Leaves soft; capitula solitary; cypselae of disk florets eglandular and fertile Microgyne

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3. Leaves rigid; capitula often in racemiform capitulescences; cypselae of disk florets glandular and sterile...... Sommerfeltia

Taxonomic treatment Podopappus tomentosus Hook. & Arn., Comp. Bot. Mag. 2: 51, 1836. Type: Brazil. Rio Grande do Sul: Asteropsis Less., Syn. Gen. Compos. 188, M. Isabelle s.n. (holotype: K?, n.v.). 1832. Neja sect. Phylloneja DC., Prodr. 5: 325, 1836. Neja macrocephala DC., Prodr. 5: 325, 1836. Type: Brazil. Rio Grande do Sul: 1833, Herbier Imperial du Brdsil 1047, Gaudichaud s.n. (lectotype, here Perennial herbs, hemicryptophytes, ligno- designated: P). tubers elongate, stems erect, basally branched or simple. Leaves densely disposed, alternate, Perennial herbs, hemicryptophytes, ligno- spirally arranged, narrowly elliptic to narrow- tubers elongate, cylindrical, 0.5-1 cm in ly obovate, sessile, apex subacute, base diameter, stems 30-45 cm tall, erect, basally attenuate, glandular and woolly-villous on branched or with only one branch, the both surfaces, margins entire. Capitula soli- branches long. Leaves densely disposed, tary (less commonly 2 or 3), terminal, sessile, alternate, spirally arranged, 15-35 x 1- heterogamous, radiate. Involucres hemispher- 3 mm, narrowly elliptic to narrowly obovate, ic to globose; phyllaries in 5 or 6 series, sessile, apex sub-acute, base attenuate, glan- glandular and villous on both surfaces. dular (vesicular biseriate trichomes, notably Receptacles epaleate. Ray florets 3- or 4- sunken on epidermis) and woolly-villous seriate, pistillate, corollas white, true rays (flagellate-septate trichomes) on both surfa- with long limbs, style branches linear. Disk ces, margins entire, the basal leaves with one florets numerous, bisexual (functionally or two pair of lobes. Capitula solitary (rarely male), corolla actinomorphic, 5-lobed; anther grouped in 2- or 3-headed capitulescences), base slightly sagittate, apical appendages terminal, sessile, heterogamous, radiate. Invo- ovate; style branches narrowly triangular to lucres 12-20 x 20-25 mm, hemispheric to oblong, totally dorsally covered by sweeping globose; phyllaries in 5 or 6 series, tips hairs. Cypselae flattened, strongly 2-nerved, reddish, outer phyllaries 7-9 x 0.7-1.1 mm, gradually attenuate towards the apex, glandu- narrowly elliptic to linear, subulate, margin lar throughout its surface and villous mainly membranous, glandular and woolly-villous on the margins, cypselae of disk florets on both surfaces; inner phyllaries 10-13 x noticeably thinner, with or without a devel- 1-1.2 mm, linear, apex subulate, margin oped ovule. Pappus of 2 or 3 series of scabrid membranous, glandular and woolly-villous bristles, sometimes with some outermost on abaxial surface, glabrous or with little bristles very short. glandular-villous pubescence towards the apex on adaxial surface. Receptacles epaleate, with short scales. Ray florets arranged in 3 or Asteropsis megapotamica (Spreng.) Mar- 4 series, 100-150, pistillate, corollas white, true chesi, Bonifacino & Sancho, comb. nov. ray, tube ca. 4 mm long, limb ca. 17 x 1 mm, Aster megapotamicus Spreng., Syst. veg. 3: linear, minutely 2- or 3-lobed; style ca. 4.5 mm 526, 1826. Type: [Brazil. Rio Grande: sine long, style branches 2-2.2 mm long, linear. Disk loc.] Spreng. herb. 1164, Sello s.n. (holo- Jflorets ca. 150, bisexual (functionally male), type: P). (Figs. 1, 2) corolla yellow, actinomorphic, 5-lobed, subcam- panulate, tube ca. 2 mm long, throat 3.5-4 mm long, tube and throat laxly glandular (biseriate Asteropsis macrocephala Less., Syn. gen Compos. glandular trichomes), lobes ca. 1 x 0.5-0.6 mm, 188, 1832. Podocoma macrocephala (Less.) Herter, triangular, glandular (vesicular biseriate glan- Florula Urug. P1. vasc. 123, 1930. Type: [Brazil.] dular trichomes) towards the apex; anthers ca. Est. Maxada, April 1823, Sello [1853?] (holotype: 2 mm long, antheropodium present ca. 0.6 mm B, destroyed; photo: F-14909; isotypes: BM, K, P). long, anther base slightly sagittate, apical

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FIG. 1. Asteropsis megapotamica. A. Habit. B. Leaf. C. Cross section of leaf showing sunken, vesicular, biseriate, glandular trichomes and flagellate-septate trichomes. D. Ray corolla with style. E. Style of ray corolla (distal part). F. Cypsela of ray floret with pappus. G. Disk corolla. H. Stamen. I. Style of disk floret (distal part). J. Cypsela of disk floret with pappus. (A from Rosengurtt B7089, MVFA; B-J from Cabrera & Zuloaga 32376, LP.)

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FIGc. 2. Asteropsis megapotamica. A. Detail of capitulum showing the woolly involucre. B. Rocky habitats of Asteropsis showing flat topped hills characteristic of northwestern Uruguay. C. Detail of the showy and radiate capitulum. D. Clumps of plants of Asteropsis, soon after re-starting vegetative growth. E. Proximal leaves. F. Distal leaves. G. Asteropsis in its habitat.

connectival appendages ca. 0.6 x 0.3 mm, hairs. Cypselae 4-4.5 mm long, elliptic, ovate, style ca. 5 mm long, style branches ca. flattened, strongly 2-nerved, gradually attenu- 2 mm long, narrowly triangular to oblong, ated towards the apex, glandular throughout its whole abaxial surface covered by sweeping surface, and villous mainly on the margins.

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Pappus ca. 7 mm long, of 2 or 3 series of from B at F shows a label with the probable scabrid, subequal bristles, sometimes with collector number, 1853. some outermost bristles very short, I mm long, all bristles tapering, somewhat flattened at the base. Additional specimens examined. BRAZIL. Rio GRANDE DO SUL:Tupareceta, 8 Mar 1939, Avila de Distribution and ecology.-Asteropsis is Araujo 28 (LP); 1833, Herbier Imp6rial du Bresil 1001, distributed in northern Uruguay (Rivera and Gaudichaud s.n. (P); 1833, Herbier Imphrial du Bresil Tacuaremb6) and southern Brazil (Rio 1006, Gaudichaud s.n. (P); Cruz Alta, 4 Dec 1893, Grande do Sul). In Uruguay, Asteropsis Malme s.n., Exped. Primae Regnellian Phanerogamae 7289 (BM); Porto Alegre, Aug 1951, Mokrdieck 2 (LP); megapotamica occurs on rocky grasslands Cruz Alta, 23 Jan 1964, Pereira & Pabst 8494, 7869 and slopes of riverbeds amid sparse shrubs (LP); without precise locality, 19 Jan 1940, Rau s.n. (SI of Baccharis spp., Eupatorium spp., Hetero- 25562). thalamus alienus (Spreng.) Kuntze, Verno- URUGUAY. RIVERA: Subida de Pena, Ruta 30, km 111.5, 11 Apr 1984, Bayce et al. s. n. (MVFA 17278); 5 nanthura nudiflora (Less.) H. Rob., and km al E de Ruta 5, Cerro Miriiiaque, 10 Apr 1984, Bayce Butia paraguariensis (Barb. Rodr.) L. H. et al. s.n. (MVFA 17226); Cerro Alegre, 12 km al S de Bailey ("dwarf butia" palms, Fig. 2B). In Tranqueras, 13 Apr 1984, Bayce et al. s.n. (MVFA certain locations, it is possible to see 17340); Cerro Hospital, 19 Dec 1901, Berro 5111 (K, abundant clumps of Asteropsis in grass-like MVFA); Ruta 30, pr6ximo a subida de Pena, 31 8' 51" S, 550 53' 53" W, 15 Jan 2006, Bonifacino 2130 vegetation dominated by the also rare and (MVFA); Entre Paso Vargas y Paso Plat6n, 15 Mar geographically very restricted Schlechtenda- 1962, Del Puerto 1747 (MVFA); Estancia Ripoll, Ruta 5, lia luzulaefolia Less. From a geomorpho- pr6ximo a Ruta 29, 28 Feb 1962, Del Puerto 1523 logic point of view, the habitat ofAsteropsis (MVFA); Valle Arroyo Platon, pr6ximo a Escuela 56, 20 Feb 1966, Marchesi 1442 (MVFA); Picada de Castro, (in northwestern Uruguay) is very interesting Escuela Agraria, 30 Jan 1958, Rosengurtt B7089 since the species grows intimately associated (MVFA); Cerro Aurora, 10 Dec 1961, Rosengurtt with flat-topped hills (Fig. 2B), a rather B8404 (MVFA). TACUAREMBO: Gruta de Los Cuervos, dramatic landscape feature of Gondwanic 8 Feb 1981, Cabrera & Zuloaga 32376 (LP, SI); origin. According to Bossi et al. (1998), the Quebrada de los Cuervos, Jan 1940, Chebataroff 10317 (LP); Gruta de los Cuervos, 9 Mar 1966, Rosengurtt et flat-topped hills ("cerros chatos") that charac- al. 10035 (MVFA, US); Gruta de Los Cuervos, 20 Feb terize northwestern Uruguay are constituted by 2004, Sancho & Bonifacino 99, 100 (LP, MVFA). sandstone belonging to Tacuaremb6 and Rivera geological formations that were formed during the Jurassic. These sandstone forma- Acknowledgments tions are composed of medium to fine-grain We thank Dan Nicolson for useful advice sand as well as some kaolinite clay elements, in nomenclature matters. We are also grateful and extend past the Uruguayan borders into to the curators of BM, K, LP, MVFA, P, SI, Brazil where Asteropsis also occurs. In Brazil, and US who provided material for examina- Asteropsis megapotamica has been found on tion. Research for this article was supported sandy soils of "campos" formations in Rio by Facultad de Agronomia (JMB, EM), Grande do Sul, an area mostly dominated by Myndel Pedersen Foundation (JMB), Smith- grasslands. sonian Institution (JMB), and Agencia Nacio- Phenology.-Asteropsis megapotamica nal de Promoci6n Cientifica y Tecnol6gica, regains activity producing branches and SECYT, Argentina (GS). leaves starting at the beginning of October and flowers and fruits from January to April. Literature Cited

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